[1] Massachusetts Inst Technol Woods Hole Oceanog Ins, Cambridge, MA 02139 USA
[2] Woods Hole Oceanog Inst, Dept Biol, Woods Hole, MA 02543 USA
[3] Univ Rhode Isl, Grad Sch Oceanog, Narragansett, RI 02882 USA
[4] Univ Rhode Isl, Dept Cell & Mol Biol, Kingston, RI 02881 USA
[5] Columbia Univ, Lamont Doherty Earth Observ, Dept Earth & Environm Sci, Palisades, NY 10964 USA